Taphonomy of modern deep,cold‐temperate water coral reefs

Deep‐water corals are widely distributed along the cold‐temperate northeastern Atlantic continental margin. Despite the widespread occurrence of these aphotic coral constructions in deep shelf settings, the processes of framework formation and postmortem alterations which result in different preservational styles are still poorly known. Detailed mapping surveys on probably one of the largest Lophelia reef structures were carried out on the Sula Ridge, Mid‐Norwegian Shelf in 270 to 300 m depth. Side scan sonar records and camera surveys yield information at various scales of resolution on the reef complex which is more than 9 km long and up to 45 m high. Living Lophelia colonies effectively prevent colonization by other organisms and are successful in the rejection of passing detrital material from the soft tissue. In a healthy condition the coral is able to encrust repetitively attached organisms by selectively secreted sclerenchyme layers, thus, this defensive reaction results in the thickening of the skeleton. Early postmortem alteration in Lophelia colonies is introduced by the formation of a biofilm and Dodgella (fungi) infestation. The biofilm is associated with selective Fe‐Mn precipitation on the coral skeleton. This is the zone of intense attachment of sessile invertebrates such as serpulids, brachiopods, foraminifers and encrusting bryozoans. More advanced taphonomic stages show an increasing dominance in sponges which reduce the interskeletal framework porosity significantly. In addition, boring sponges excavate the thickly calcified Lophelia skeletons, thus leading to in situ collapsing structures on the sea floor. It is the intensity of sediment trapping biofilms and sponge colonization and the amount of imported detrital particles predominantly from the pelagial zone that control the generation of a pure coral rubble facies or the preservation of collapsed but mud‐rich detrital mounds.     

Comparison of deep-water coral reefs and lithoherms off southeastern USA

Two types of deep-water coral bioherms occur off the coast of southeastern United States: Oculina and Lophelia/Enallopsammia. The deep-water Oculina bioherms form an extensive reef system at depths of 70–100 m along the shelf edge off central eastern Florida. These reefs are comprised of numerous pinnacles and ridges, 3–35 m in height. Each pinnacle is a bank of unconsolidated sediment and coral debris that is capped on the slopes and crest with living and dead colonies of Oculina varicosa, the ivory tree coral. In comparison, deep-water reefs of Lophelia pertusa and Enallopsammia profunda corals occur at depths of 500–850 m (maximum 150-m relief) along the base of the Florida-Hatteras slope in the Straits of Florida. On the western edge of the Blake Plateau off South Carolina and Georgia, 54-m high banks of Enallopsammia and Lophelia occur at depths of 490–550 m, whereas on the eastern edge of the plateau the reefs form structures 146 m in height and at depths of 640–869 m. The geomorphology and functional structure of both the Oculina and Lophelia reefs are similar. North of Little Bahama Bank, at depths of 1000–1300 m, a region of bioherms is dominated by the coral Solenosmilia sp.; Lophelia is reportedly absent. This paper summarizes 25 years of submersible studies on the deep-water Oculina reefs, describes submersible reconnaissance of deep-water Lophelia reefs off the southeastern United States, and contrasts these types of bioherms with the deep-water lithoherms in the Straits of Florida west of the Bahamas.     

Shallow-buried Pleistocene Madrepora-dominated coral mounds on a muddy continental slope, Tuscan Archipelago, NE Tyrrhenian Sea

Subfossil azoxanthellate deep-sea coral mounds occur at 355–410 m on the continental slope of the NE Tyrrhenian Sea between Gorgona and Capraia islands, Tuscan Archipelago. These low-relief patch reefs are at present buried by a thin muddy drape. Their age is latest Pleistocene. The colonial scleractinian Madrepora oculata is the major frame builder, in association with the solitary coral Desmophyllum dianthus and the colonial coral Lophelia pertusa. These NE Tyrrhenian Madrepora-dominated coral mounds represent one of the few known Mediterranean examples of deep-coral colonization of a muddy, low-gradient continental slope.     

Coral reefs in the Southern Barents Sea: habitat description and the effects of bottom fishing

Fifteen Lophelia reefs from offshore to coastal areas off northern Norway were studied using video. Health status of the coral habitat (degree of physical impact, % cover of living tissue, colony size), occurrence of trawl marks and lost fishing gear, height of coral colonies and associated fauna were analysed from 44 video-lines. Fishing impact was more frequent on the offshore reefs (36.5% of the observed areas) than those in the coastal reefs (0.6%). The most visible effects of fishing were broken and displaced coral colonies. At some sites only small scattered fragments of live corals were observed, indicating recent impact. The mean colony height of Lophelia and gorgonian corals at impacted sites was around half the size of those at non-impacted sites. Both species richness and abundance was higher at non-impacted coral habitats compared to impacted. The actinarian Protanthea simplex and unidentified brittlestars were the only taxa with higher abundance on impacted compared with non-impacted habitats. The reefs at the offshore location were protected against bottom trawling in 2009 through the establishment of a marine protected area (MPA), but a general ban against trawling on known coral reefs had already been implemented in 1999. In the MPA, signs of regrowth were observed. Most of the observed damage probably occurred over 10 years earlier. Results show that live and non-impacted cold water coral reefs have an important ecological function by enhancing the local biodiversity and fish abundance. Preventing further damage to impacted reefs may lead to full recovery within a few decades.     

Spawning of the colonial coral <Emphasis Type="Italic">Cladocora caespitosa</Emphasis> (Anthozoa,Scleractinia) in the Southern Adriatic Sea

Data on sexual reproduction of scleractinian coral species living in temperate zones, particularly in the Mediterranean Sea, are quite scarce. This paper describes sexual reproduction of the colonial coral Cladocora caespitosa from Veliko jezero (Mljet Island) in the Adriatic Sea. Spawned orange eggs and white sperm bundles were observed on the coral bank of C. caespitosa two nights before the full moon (20 June 2005) coinciding with increasing water temperature and correlated with the lunar cycle. Spawning was observed during five nights, involving about 30% of the colonies from the coral bank. Different colonies on the bank released only one type of gamete during the reproductive period. The diameter of the sperm bundles ranged from 100 to 200 μm (average 163 μm; SD = 47.08), while the female gametes diameter ranged from 300 to 500 μm (average 416 μm; SD = 73.12).     

Coral associations of the Pleistocene Ironshore Formation,Grand Cayman

The 125-ka sea level, which was approximately 6 m above present-day sea level, led to the partial flooding of many Caribbean islands. On Grand. Cayman, this event led to the formation of the large Ironshore Lagoon that covered most of the western half of the island and numerous, small embayments along the south, east, and north coasts. At that time, at least 33 coral species grew in waters around Grand Cayman. This fauna, like the modern coral fauna of Grand Cayman, was dominated byMontastrea annularis, Porites porites, Acropora polmata, andA. cervicornis. Scolymia cubensis andMycetophyllia ferox, not previously identified from the Late Pleistocene, are found in the Pleistocene patch reefs.Madracis mirabilis, Colpophyllia breviserialis, Agaricia tenuifolia, A. lamarcki, A. undata, Millepora spp., Mycetophyllia reesi, M. aliciae, andM. danaana, found on modern reefs, have not been identified from the Late Pleistocene reefs. Conversely,Pocillopora sp. cf.P. palmata, which is found in Late Pleistocene reefs, is absent on the modern reefs around Grand Cayman. The corals in the Ironshore Formation of Grand Cayman have been divided into 10 associations according to their dominant species, overall composition, and faunal diversity. Many of these associations are similar to the modern associations around Grand Cayman. Each of the Pleistocene coral associations, which can be accurately located on the known Late Pleistocene paleogeography of Grand Cayman, developed in distinct environmental settings. Overall trends identified in the modern settings are also apparent in the Late Pleistocene faunas. Thus, the diversity of the coral faunas increased from the interior of the Ironshore Lagoon to the reef crest. Similarly, the coral diversity in the Pleistocene patch reefs was related to the size of the reefs and their position relative to breaks in the barrier reef. The barrier reef included corals that are incapable of sediment rejection; whereas the patch reefs lacked such corals.     

Recent benthic foraminiferal assemblages from cold-water coral mounds in the Porcupine Seabight

Cold-water coral ecosystems are characterised by a high diversity and population density. Living and dead foraminiferal assemblages from 20 surface sediment samples from Galway and Propeller Mounds were analysed to describe the distribution patterns of benthic foraminifera on coral mounds in relation to different sedimentary facies. Hard substrates were examined to assess the foraminiferal microhabitats and diversities in the coral framework. We recognised 131 different species, of which 27 prefer an attached lifestyle. Epibenthic species are the main constituents of the living and dead foraminiferal assemblages. The frequent species Discanomalina coronata was associated with coral rubble, Cibicides refulgens showed preference to the off-mound sand veneer, and Uvigerina mediterranea displayed abundance maxima in the main depositional area on the southern flank of Galway Mound, and in the muds around Propeller Mound. The distribution of these species is rather governed by their specific ecological demands and microhabitat availability than by the sedimentary facies. Benthic foraminiferal assemblages from coral mounds fit well into basin-wide-scale distribution patterns of species along the western European continental margin. The diversity of the foraminiferal faunas is not higher on the carbonate mounds as in their vicinity. The living assemblages show a broad mid-slope diversity maximum between 500 and 1,300 m water depth, which is the depth interval of coral mound formation at the Celtic and Amorican Margin. The foraminiferal diversity maximum is about 700 m shallower than comparable maxima of nematodes and bivalves. This suggests that different processes are driving the foraminiferal and metazoan diversity patterns.     

Brown-band syndrome on feeding scars of the crown-of-thorn starfish Acanthaster?planci

Despite the growing impact of coral diseases on reef ecosystems, little is known about the role of coral predation in disease transmission. An experiment on the coral reefs of Derawan Island, Indonesia, revealed brown-band syndrome on Acropora cytherea coral colonies following predation by the crown-of-thorn starfish Acanthaster planci. To experimentally exclude predation, living coral tissue adjacent to feeding scars was enclosed using cages and monitored for 15 days. Compared with similarly caged but uninjured colonies, which showed no sign of disease or tissue loss, preyed upon colonies showed a higher incidence of the disease, coupled with further tissue mortality. This study provides preliminary evidence that A. planci might promote the transmission of some coral diseases.     

Comparing Deep-Sea Fish Fauna between Coral and Non-Coral “Megahabitats” in the Santa Maria di Leuca Cold-Water Coral Province (Mediterranean Sea)

Two experimental longline surveys were carried out in the Santa Maria di Leuca (SML) cold-water coral province (Mediterranean Sea) during May–June and September–October 2010 to investigate the effect of corals on fish assemblages. Two types of “megahabitat” characterized by the virtual absence of fishing were explored. One was characterized by complex topography including mesohabitats with carbonate mounds and corals. The other type of megahabitat, although characterized by complex topographic features, lacks carbonate mounds and corals. The fishing vessel was equipped with a 3,000 m monofilament longline with 500 hooks and snoods of 2.5 m in length. A total of 9 hauls, using about 4,500 hooks, were carried out both in the coral megahabitat and in the non-coral megahabitat during each survey. The fish Leucoraja fullonica and Pteroplatytrygon violacea represent new records for the SML coral province. The coral by-catch was only obtained in the coral megahabitat in about 55% of the stations investigated in both surveys. The total catches and the abundance indices of several species were comparable between the two habitat typologies. The species contributing most to the dissimilarity between the two megahabitat fish assemblages were Pagellus bogaraveo, Galeus melastomus, Etmopterus spinax and Helicolenus dactylopterus for density and P. bogaraveo, Conger conger, Polyprion americanus and G. melastomus for biomass. P. bogaraveo was exclusively collected in the coral megahabitat, whereas C. conger, H. dactylopterus and P. americanus were found with greater abundance in the coral than in the non-coral megahabitat. Differences in the sizes between the two megahabitats were detected in E. spinax, G. melastomus, C. conger and H. dactylopterus. Although these differences most probably related to the presence-absence of corals, both megahabitats investigated play the role of attraction-refuge for deep-sea fish fauna, confirming the important role of the whole SML coral province as a refuge area from fishing.     

Physiology and taxonomy of scleractinian corals: a case study in the genus Stylophora

The morphology and skeletal characteristics of colonies of the coral genus Stylophora living on the reef edge at 1 m depth on the Jordanian coast of the Gulf of Aqaba (Red Sea) are those of S. mordax (Dana 1846) which has not been reported previously from that area. These colonies were considered earlier as ecomorphs of S. pistillata (Esper 1797) which lives down to at least 67 m on the reef slopes. Growth, organic content and metabolism were compared in colonies living at different depths (1,5,10 and 30 m). The trends of twelve parameters between 1 and 5 m were different from the variation observed between 5 and 30 m. Colonies living at 1 m have a higher chlorophyll content but a lower metabolic activity and growth rate than colonies living at 5 m. Most of these pecularities cannot be explained by the influence of environmental factors. It is therefore suggested that S. mordax is a valid taxon.     

Character release following extinction in a Caribbean reef coral species complex

Abstract The Pleistocene extinction of the widespread organ‐pipe Montastraea coral had measurable morphological and ecological effects on surviving lineages of the Montastraea“annularis” species complex. Extinction of the organ‐pipe Montastraea occurred after more than 500,000 years of dominance in the shallow‐water reef habitat of Barbados. Extinction resulted in a morphological shift of the columnar Montastraea lineage from thick to thin columns in modern reef environments. Pleistocene colonies of the columnar morphotype sympatric with organ‐pipe Montastraea showed greater column widths than those in allopatry. We subjected our data to a number of criteria for interpreting the morphological shift as character release following lifting of competitive pressure after extinction. The morphological differences do not appear to be due either to chance or to physical properties of the marine environment. Differential local extinction and recolonization of four members of the species complex did not occur on Barbados, so that the species coexisted and appear to have coevolved between more than 600,000 and 82,000 years ago. The morphological shift is related to coral growth form and growth rate, and thus reflects the acquisition of a primary resource in corals‐light. Character release occurred at the same oceanic Caribbean island (Barbados) where environments have fluctuated with similar variance throughout the period of coexistence. Not only has competition among living members of the Montastraea“annularis” species complex been convincingly demonstrated, but trends in relative abundance among fossil members of the species complex strongly suggest that a competitive hierarchy was operating during their Pleistocene coexistence on Barbados. We also observed an ecological analogue to character release on another Caribbean island, Curaçao. The distribution and abundance of living columnarM. annularis s.s. and massive M. faveolata from the leeward reef crest in Curaçao is greater now than in the Pleistocene, when organ‐pipe Montastraea dominated this shallow‐water reef habitat. Extinction of the faster growing, shallow‐water organ‐pipe Montastraea resulted in higher abundance of the columnar Montastraea lineage in shallow‐water habitats, where it shifted its morphology to one adapted to high light levels. The species extinction released surviving lineages from a competitive network that had resulted in lower rank abundance in the Pleistocene community and enhanced abundance of both columnar M. annularis s.s. and M. faveolata in modern communities. Full validation of our interpretation of character release must await experiments that demonstrate whether phenotypic differences between populations have a genetic basis. However, we believe the results of this study point to the important, yet heretofore neglected, role that biological interactions have played in the evolution of closely related reef coral species.     

Morphology and sedimentology of (clustered) cold-water coral mounds at the south Rockall Trough margins,NE Atlantic Ocean

Cold-water coral mounds on both margins of the Rockall Trough (NE Atlantic Ocean) have a strongly different morphology. Single, isolated mounds occur on the SE margin and are mainly found on the upper slope between 900 and 650 m water depth, while large mound clusters are found on the SW margin in water depths between 600 and 1,000 m, in a narrow zone almost parallel to the slope. Sedimentation rates on the mounds are higher than on the surrounding seabed as a result of baffling of biogenic carbonate debris and siliciclastic particles by the coral framework covering the mounds. This is confirmed by ²¹⁰Pb measurements. The individual coral growth rate can be three times higher then the vertical growth rate of the coral cover (±10 mm year⁻¹) which in turn is more than an order of magnitude higher then the present-day overall mound growth rate (±0.25 mm year⁻¹). The presence of extensive hardgrounds and firmgrounds and the three-dimensional coral framework are considered to be responsible for the stability of the relatively steep slopes of the mounds. High current velocities in the intramound areas result in local non-sedimentation and erosion, as is shown by the presence of IRD (ice-rafted debris) lag deposits on the seabed and moats around some of the mounds. The morphology and sedimentology of cold-water coral-covered (mainly Lophelia pertusa and Madrepora oculata) mounds on the southern Rockall Trough margins (NE Atlantic Ocean) is discussed and a model describing the development of these mounds is presented.     

Mapping Cold-Water Coral Habitats at Different Scales within the Northern Ionian Sea (Central Mediterranean): An Assessment of Coral Coverage and Associated Vulnerability

In this study, we mapped the distribution of Cold-Water Coral (CWC) habitats on the northern Ionian Margin (Mediterranean Sea), with an emphasis on assessing coral coverage at various spatial scales over an area of 2,000 km² between 120 and 1,400 m of water depth. Our work made use of a set of data obtained from ship-based research surveys. Multi-scale seafloor mapping data, video inspections, and previous results from sediment samples were integrated and analyzed using Geographic Information System (GIS)-based tools. Results obtained from the application of spatial and textural analytical techniques to acoustic meso-scale maps (i.e. a Digital Terrain Model (DTM) of the seafloor at a 40 m grid cell size and associated terrain parameters) and large-scale maps (i.e. Side-Scan Sonar (SSS) mosaics of 1 m in resolution ground-truthed using underwater video observations) were integrated and revealed that, at the meso-scale level, the main morphological pattern (i.e. the aggregation of mound-like features) associated with CWC habitat occurrences was widespread over a total area of 600 km². Single coral mounds were isolated from the DTM and represented the geomorphic proxies used to model coral distributions within the investigated area. Coral mounds spanned a total area of 68 km² where different coral facies (characterized using video analyses and mapped on SSS mosaics) represent the dominant macro-habitat. We also mapped and classified anthropogenic threats that were identifiable within the examined videos, and, here, discuss their relationship to the mapped distribution of coral habitats and mounds. The combined results (from multi-scale habitat mapping and observations of the distribution of anthropogenic threats) provide the first quantitative assessment of CWC coverage for a Mediterranean province and document the relevant role of seafloor geomorphology in influencing habitat vulnerability to different types of human pressures.     

Coral mass- and split-spawning at a coastal and an offshore Venezuelan reefs,southern Caribbean

This study aimed to evaluate potential differences in coral spawning behavior between a fringing coastal reef and an offshore reef in the southern Caribbean. For this, scleractinian and gorgonian colonies (N = 324) of 21 species were mapped along eight transects, each 10-m long, at two study sites located in Morrocoy and Los Roques National Parks, Venezuela. Observations were made between 19:30 and 23:00 from August 23 to 30 and from September 26 to 30, 2002. Ninety one colonies belonging to six hard coral and seven octocoral species spawned or planulated during this period. We were unable to observe any signs of reproductive activity in 95 colonies of nine species different from those that reproduced. Despite the differences in environmental conditions between the two sites, we observed high synchrony in the spawning behavior of seven coral species common to both reefs. The most striking result was the ability of colonies of Montastraea faveolata and Eusmilia fastigiata to split spawn up to three times, either in consecutive nights or in different months.     

Spatial Homogeneity of Bacterial Communities Associated with the Surface Mucus Layer of the Reef-Building Coral Acropora palmata

Coral surface mucus layer (SML) microbiota are critical components of the coral holobiont and play important roles in nutrient cycling and defense against pathogens. We sequenced 16S rRNA amplicons to examine the structure of the SML microbiome within and between colonies of the threatened Caribbean reef-building coral Acropora palmata in the Florida Keys. Samples were taken from three spatially distinct colony regions—uppermost (high irradiance), underside (low irradiance), and the colony base—representing microhabitats that vary in irradiance and water flow. Phylogenetic diversity (PD) values of coral SML bacteria communities were greater than surrounding seawater and lower than adjacent sediment. Bacterial diversity and community composition was consistent among the three microhabitats. Cyanobacteria, Bacteroidetes, Alphaproteobacteria, and Proteobacteria, respectively were the most abundant phyla represented in the samples. This is the first time spatial variability of the surface mucus layer of A. palmata has been studied. Homogeneity in the microbiome of A. palmata contrasts with SML heterogeneity found in other Caribbean corals. These findings suggest that, during non-stressful conditions, host regulation of SML microbiota may override diverse physiochemical influences induced by the topographical complexity of A. palmata. Documenting the spatial distribution of SML microbes is essential to understanding the functional roles these microorganisms play in coral health and adaptability to environmental perturbations.     

Depth limit for reef building corals in the Au’au Channel, S.E. Hawaii

In this paper, the relationship between reef building (accretion) and depth in an optimal inter-island channel environment in Hawaii is analyzed. For accretion, the growth rate of Porites lobata is used as a proxy for the reef community, because it is the most abundant and dominant species of reef building coral in Hawaii. Optimal growth of P. lobata occurs at a depth of 6 m, below which both growth rate and abundance decrease with increasing depth. A lower depth limit for this species is found at about 80–100 m, yet reef accretion ceases at ~50 m depth. Below 50 m, rates of bio-erosion of colony holdfasts equal or exceed the growth of basal attachments, causing colonies to detach from the bottom. Continued bio-erosion further erodes and dislodges colonies leading to their breakdown and ultimately to the formation of coralline rubble and sand. Thus, within this channel environment in Hawaii, a threshold for reef building exists at ~ 50 m depth, where coral accretion is interrupted by bio-erosion. Conceptually viewed, this depth horizon is analogous to a vertical Darwin Point, although quite narrow in space and time. More importantly, it explains the history of reef morphology in the Au’au Channel where a chronological hiatus exists at a depth near 50 m. This hiatus separates shallower modern growth (about 100 years or less) from the deeper reef which is all due to accretion during the early Holocene or Pleistocene epochs.     

Distribution and abundance of fish in deep-sea coral habitats

Experimental fishing with long-lines and gillnets was conducted on the continental shelf off southwestern Norway between 150 and 350 m depth. Abundance and distribution of redfish (Sebastes marinus L., 1758), ling (Molva molva L., 1758), and tusk (Brosme brosme Ascanius, 1772) were quantified in Lophelia pertusa (L., 1758) coral reefs and in non-coral habitats. The largest catches of redfish were made with long-line fleets set in coral reef habitats. Ling and tusk were also most numerous in coral habitats, although not statistically significant. Fish caught in coral habitats tended to be larger in size than in non-coral habitats. The diet of redfish, tusk and ling included the same prey groups in all habitats, but they differed at the species level. Lophelia-reefs may provide a profitable feeding place for tusk. For the planktivorous Sebastes, on the other hand, their affinity to the reefs seems primarily to be related to the physical structure offered by the reefs.     

Growth of Mediterranean reef of<Emphasis Type="Italic"> Cladocora caespitosa</Emphasis> (L.) in the Late Quaternary and climate inferences

A sclerochronological analysis was performed on Cladocora caespitosa corals from Late Pleistocene terraces near Taranto (Apulia, Italy) to reconstruct the main palaeoenvironmental conditions at the time of their growth. The fossil corallites were sampled in the Santa Teresiola uplifted bank or ‘open frame reef’ attributed to the Last Interglacial Period. The typical, annual growth pattern of the temperate coral with two alternate high- and low-density bands allowed the reconstruction of two multidecadal growth curves of 61 and 95 years. Trend analysis showed oscillations in annual growth rates similar to those observed in recent, living colonies sampled along a north–south latitudinal transect around the Italian and Croatian coasts as far as Tunisia. The mean growth rate of the fossil reef (4.2 ± 2 mm year⁻¹) is comparable to those measured on colonies living in the coldest part of the Mediterranean Sea. The comparison with data from living Croatian banks shows how fossil C. caespitosa lived in a semi-enclosed environment characterized by seasonal inputs of fresh, cold water. The greatest variations in decadal growth rates of the fossil colonies support the hypothesis of larger amplitude of the seasonal cycles in the past. The death of the fossil bank was probably due to a sudden alluvial input that suffocated the reef with a great amount of mud. Another possible cause of the death of the bank was a prolonged increase in summer temperatures that caused colony mortality and enhanced algal colonization.     

The distribution,abundance, and the effects of fire on mound building termites (Trinervitermes and Cubitermes spp., Isoptera: Termitidae) in northern guinea savanna West Africa

Summary Termite mound densities in typical guinea savanna, Detarium, and grassland (boval) habitats in northern guinea savanna were determined by random quadratting of 2–3 sites in each habitat (100, 10x10 m quadrats per habitat). Dominant species in guinea savanna were T. geminatus (46 mounds ha^-1) and T. oeconomus (21 mounds ha^-1), in Detarium T. geminatus (59 mounds ha^-1) and C. curtatus (45 mounds ha^-1) and in boval C. curtatus (72 mounds ha^-1) and T. geminatus (22 mounds ha^-1). Only C. curtatus densities and total densities differed significantly between sites within habitats, but all species differed significantly in abundance between habitats. The composition of each community was related to general environment but no particular environmental variable was shown to be a major determinant of termite distribution. Evidence for the limitation of termite populations was obtained from indirect evidence of competition between colonies in Detarium, and by experimental manipulation of fire regimes in the typical guinea savanna habitat. Harvester termites increased four-five fold over two years in fire-protected plots as a result of increased food supplies. Total termite densities in the fire-protected community equilibrated to the new population density (100 mounds ha^-1) after only two-three years.     

Substrate pitting and boring pattern of Hyrrokkin sarcophaga Cedhagen, 1994 (Foraminifera) in a modern deep-water coral reef mound

Colonization structures of the large parasitic foraminifer Hyrrokkin sarcophaga Cedhagen, 1994 on Lophelia pertusa, Acesta excavata and Delectopecten vitreus are described from a deep-water coral reef mound on the mid-Norwegian shelf at 240 to 300 m water depth. Hyrrokkin sarcophaga is the only epibiont which is capable of attaching itself on the soft tissue-protected coral skeleton where it tends to form clusters of 3 to 8 specimens close to the tentacles of Lophelia. The foraminifer excavates a pit up to 1.5 mm deep and etches a straight channel through the skeleton of the host which ends within the soft tissue. In contrast to Lophelia, infested bivalves show a strong wound repair reaction and seal the etched channels by intense calcification. The etching is only performed by adult specimens. Substrate pitting is considered to improve the attachment strength while boring enables the parasite to secure a persistent nutrient source.